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Bat Activity and Use of Hibernacula in Wood Buffalo National Park, Author(s): Jesika P ReimerCori L LausenRobert MR BarclaySharon Irwin, Mike K Vassal Source: Northwestern Naturalist, 95(3):277-288. 2014. Published By: Society for Northwestern Vertebrate Biology DOI: http://dx.doi.org/10.1898/13-30.1 URL: http://www.bioone.org/doi/full/10.1898/13-30.1

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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. NORTHWESTERN NATURALIST 95:277–288 WINTER 2014

BAT ACTIVITY AND USE OF HIBERNACULA IN WOOD BUFFALO NATIONAL PARK, ALBERTA

JESIKA PREIMER Department of Biological Sciences, University of , 2500 University Drive NW, Calgary, AB T2N 1N4 ; [email protected]

CORI LLAUSEN Wildlife Conservation Society Canada, PO Box 606, Kaslo, BC V0G 1M0 Canada

ROBERT MR BARCLAY Department of Biological Sciences, University of Calgary, 2500 University Drive NW, Calgary, AB T2N 1N4 Canada

SHARON IRWIN,MIKE KVASSAL Wood Buffalo National Park, Parks Canada, PO Box 750, Fort Smith, NT X0E 0P0 Canada

ABSTRACT—Relatively little is known about bats or bat hibernacula in . We were interested in documenting species diversity and seasonal activity of bats in Wood Buffalo National Park, including use of a cave hibernaculum by Little Brown Myotis (Myotis lucifugus), Northern Myotis (Myotis septentrionalis), and Big Brown Bats (Eptesicus fuscus). We used acoustic monitoring and mist netting over 3 y to assess species diversity and seasonal activity. We also recorded cave temperature during hibernation. During the summers of 2010 to 2012, we captured 470 bats including M. lucifugus, M. septentrionalis, and E. fuscus. We identified 2 migratory species via echolocation recordings in 2011: Lasiurus cinereus (Hoary Bat) was recorded in the area from mid-May to early October, and Lasiurus borealis (Eastern Red Bat) from mid-June to early August. Resident bat activity at the hibernaculum was greatest from mid-June to early September. Our findings provide a first approximation of species diversity and describe seasonal activity patterns of bats in Wood Buffalo National Park.

Key words: Alberta, Big Brown Bat, Eastern Red Bat, Eptesicus fuscus, hibernaculum, Hoary Bat, Lasiurus borealis, Lasiurus cinereus, Little Brown Myotis, Myotis lucifugus, Myotis septentrionalis, Northern Myotis, seasonal bat activity, Wood Buffalo National Park

Species diversity and seasonal activity of bats understand the complexity of local ecosystems, have been well documented throughout much provides baseline data in an area already of North America. While initial studies on bats impacted by climate change (for example, in northern Canada have been focused in Payette and others 2004), and provides insight southern (Jung and others 2006; Slough into how species behave at the northern limit of and Jung 2008; Talerico 2008), southwestern their ranges. In addition, with the spread of (Lausen and others white-nose syndrome (WNS; a fungal disease 2014), and the Fort MacMurray area of north- killing millions of bats in North America; eastern Alberta (Grindal and others 2011), there Blehert and others 2009), investigating condi- remains a large portion of northern Canada tions in bat hibernacula at northern latitudes where we know relatively little about bats, may provide insight into the survivability of including large, undeveloped areas such as Pseudogymnoascus destructans (fungus that caus- Wood Buffalo National Park (WBNP). Investi- es WNS; Minnis and Lindner 2013) in these gating the presence and seasonal activity of bat potentially colder environments. Therefore, the species in these remote areas helps us better goals of this study were to survey bat species

277 278 NORTHWESTERN NATURALIST 95(3) diversity in a portion of the park, describe secondary entrance as the ‘ice chute’. Surround- seasonal activity of resident and migratory ing trees shade both entrances and standing bats, and investigate microclimatic conditions water in the sinkholes remains frozen for at one of the most northern bat hibernacula 9 months of the year. The cave is approximately currently known (Walk-in Cave), located in 1 km long and has 4 main sections: (1) the northeastern Alberta, Canada. entrance hall (approximately 2–5-m high, 6-m Based on previous bat surveys in northern wide, and 90–100-m long); (2) the middle Alberta and the Northwest Territories (van Zyll chamber (approximately 6-m high, 30–45-m de Jong 1985; Caceres and Pybus 1997; Grindal wide, and 90-m long); (3) the main chamber and others 2011; Reimer 2013; Wilson and (15-m high, 20-m wide, 120–180-m long); and (4) others 2014), we expected Little Brown Myotis the ice chute chamber near the secondary (Myotis lucifugus), Northern Myotis (M. septen- entrance of the cave. An additional small back trionalis), and Big Brown Bats (Eptesicus fuscus) chamber (approximately 10-m deep, 1.5-m high, to be present in Wood Buffalo National Park. and 2-m wide) holds the majority of the Given recent observations south (Grindal and hibernating bats (Lewis 1974–1975; hereafter, others 2011) and west (Nagorsen and Paterson the ‘‘Bat Chamber’’). Two additional side cham- 2012; Lausen and others 2014) of our study area, bers have recently been described as: (1) the we also expected migratory species including rafting lake and aven (a side chamber and stream Hoary Bats (Lasiurus cinereus), Eastern Red Bats loop off of the ice chute chamber); and (2) the (L. borealis), and Silver-haired Bats (Lasionycteris Lewis Rink (a large side-loop passage off of the noctivagans) to be present. entrance hall; Greg Horne, pers. comm.). Based on studies at more southern latitudes (Twente 1955; Beer and Richards 1956; Hen- Acoustic Monitoring shaw and Folk 1966; Brack and Twente 1985; Davis and others 1999), hibernating bats prefer To assess species diversity and seasonal bat cave temperatures of 2.6 to 86C. We predicted activity, we recorded echolocation calls using that Walk-in Cave would provide similar AnaBat II ultrasonic detectors with CF- temperature profiles as those for hibernacula ZCAIMs set to a division ratio of 16 (Titley selected by bats at more southern locations. We Electronics, Ballina, Australia) outside Walk-in also investigated another, nearby cave (Peace Cave during 21 April 2011 to 7 October 2011 Point Cave) as a potential hibernaculum in the and 20 April 2012 to 26 October 2012; and at a southern region of WBNP. pond 3 km to the west of the cave (hereafter, WB Pond), for the period of 11 May 2011 to 8 METHODS October 2011. At Walk-in Cave, we placed a detector in a weatherproof box 3 m from the Study Site entrance and pointed the microphone away We studied bats in the Alberta portion of from the entrance to prevent recording bats WBNP, mainly in the vicinity of Walk-in Cave, flying within the cave. At WB Pond, we the largest cave currently known in the park mounted a detector on a tree trunk 2.5 m off (Figure 1). WBNP is within the Boreal Plains the ground and 1 m away from the water’s Ecozone (Ecological Stratification Working Group edge. Each detector remained in a fixed 1996), and the area is primarily boreal forest location for the duration of the survey. dominated by Trembling Aspen (Populus tremu- We reviewed all AnaBat recordings using loides) and Jack Pine (Pinus banksiana), inter- AnalookW version 3.8v. We identified species spersed with wetlands, meandering streams, and as Eptesicus fuscus, Lasiurus borealis, L. cinereus, a large number of karst landforms. Walk-in Cave Lasionycteris noctivagans, and Myotis spp., based is a limestone cave formed as an underground on minimum frequency, call duration, and slope drainage system. It has 2 entrances each located as outlined in Lausen and others (2014). Call at the bottom of large sinkholes (approximately characteristics of M. lucifugus and M. septentrio- 7-m deep). The main entrance is approximately nalis overlap (Fenton and Bell 1981), often 12-m wide by 4-m high; the secondary entrance making them indistinguishable with AnaBat II is slightly narrower and lined with ice year detectors; therefore, we grouped Myotis spp. in round. Lewis (1974–1975) referred to this the analysis. WINTER 2014 REIMER AND OTHERS:BAT ACTIVITY IN WOOD BUFFALO NP 279

FIGURE 1. Location of the study area in Wood Buffalo National Park, Alberta, Canada. Inset map is of the location of Wood Buffalo National Park within western Canada.

Bat Captures and Handling set a 10-m long 3 2.6-m high net across the mouth of the cave, approximately 1 m in front To complement our acoustic survey, we used of the entrance. At WB Pond we set 2 nets (6 m mist nets (Avinet Inc., Dryden, NY, USA) to 3 2.6 m); 1 strung across a road and the other capture bats in 2010–2012. Capture sites includ- across an opening in the forest leading to the ed: (1) the mouth of Walk-in Cave (n 5 7 pond. Nets were open from 1 to 3 h depending nights); (2) WB Pond (n 5 9); and (3) at Pine on the capture rate and night length. Upon Lake, 25 km to the southeast of the cave (n 5 1). capture, we held bats in a cloth bag for 1 h to We also investigated another known cave in allow defecation prior to weighing. We identi- WBNP (Peace Point Cave; near Peace Point, fied individuals to species based on morpho- Fig. 1). Net size and location remained consis- logical features and measurements, using the tent at sites among nights. At Walk-in Cave, we key provided by Nagorsen (2002); determined 280 NORTHWESTERN NATURALIST 95(3) sex, age (adult or juvenile), and reproductive significant positive effect of ambient tempera- condition, weighed them with a portable bal- ture at sunset on echolocation activity (linear 2 ance, measured forearm and ear lengths with regression, F1, 280 5 50.3, P , 0.001, R 5 0.15), calipers, fitted them with a uniquely labeled with greater activity occurring during warmer band (Porzana Ltd., East Sussex, UK) for future nights. identification, and released them at their point We captured M. lucifugus, M. septentrionalis, of capture. We distinguished M. lucifugus from and E. fuscus emerging from Walk-in-Cave M. septentrionalis based on ear and tragus length during spring (May to early June; Figure 3). (Nagorsen 2002). We identified volant juveniles We captured no M. septentrionalis at the cave by their evenly tapered phalangeal-metacarpal after 16 May. Fifty Myotis spp. were captured at joints and unfused epiphyseal plates observed the entrance to Walk-in Cave, of which only 2 by back-illuminating the wing membrane (Kunz were juvenile M. lucifugus. Of the 3 E. fuscus and Parsons 2009). captured in a mist net at the entrance to Walk-in In September 2010 and November 2011, we Cave on 11 September 2010, 2 were exiting surveyed bats in easily accessible chambers of within 2 h of dusk. All E. fuscus captured in Walk-in Cave, including the front, middle, main, mid- to late August, including juveniles (n 5 and Bat Chambers, to estimate the number of 24), were entering the cave 1 to 2 h after the 1st individuals. We also handled a subsample to Myotis spp. was observed exiting the cave. The determine species, sex, and age. In September majority of Myotis spp. captured exiting the 2010, we surveyed bats in Peace Point Cave. cave had mud on their bodies or wings; no mud was observed on E. fuscus captured entering the Cave Microclimate cave. To describe the temperature at Walk-in Cave, Internal Cave Inspections we recorded cave chamber temperature and ambient temperature at WB Pond, using tem- On 10 and 11 September 2010, we surveyed perature loggers (Hobo model UA-002-08; On- Walk-in Cave and observed approximately 35 set, Bourne, MA) with an accuracy of ±0.476C. Myotis spp. roosting on the walls, mainly in the We placed a temperature logger in both the Bat Chamber. We handled 27 bats, all of which middle chamber and Bat chamber of Walk-in were M. lucifugus; 21 were reproductive adult Cave. We tested the effect of ambient temper- males (swollen epidiymis clearly visible), 1 was ature on bat activity at Walk-in Cave and WB a non-reproductive adult male, 2 were young- Pond with linear regression, including bat of-the-year males, and 3 were post-lactating activity (number of recorded AnaBat files) and adult females. On 10 November 2011, we ambient temperature as dependent and inde- observed 73 Myotis spp. in all chambers of pendent variables, respectively. Due to gaps in Walk-in Cave except the entrance hall, middle outside air temperature recordings at WB Pond, chamber, and ice chute chamber (the outermost ambient temperatures for statistical analysis sections of the cave), including 22 bats in the were obtained from the Environment Canada main chamber, 41 bats in the Bat Chamber, and website (http://climate.weather.gc.ca) for the 10 distributed in various other locations. It is Fort Smith airport weather station, approximately likely that more bats were present, but in 30-km north of the study area. All statistical locations where they were not visible. We analyses were performed with Stata/SE 12.1 handled 5 M. lucifugus randomly sampled from (StataCorp, Texas). the cave wall, and all were adult males. We observed M. lucifugus using the cave as a RESULTS summer day roost (roosting in the front 3 chambers during the day and captured in mist Bat Activity at Walk-in-Cave nets as they emerged from the cave at dusk) on We recorded echolocation calls at the mouth all summer visits to the cave (15 visits from 12 of Walk-in Cave from 21 April to 22 September May to 6 September 2011 and 4 June to 13 2011 and 20 April to 21 October 2012 (Figure 2). August 2012). Specifically, we counted 30 and The highest activity for both years was recorded 47 Myotis spp. day-roosting in the front 3 during early September and there was a chambers (entrance hall, middle chamber, main WINTER 2014 REIMER AND OTHERS:BAT ACTIVITY IN WOOD BUFFALO NP 281

FIGURE 2. Bat echolocation activity recorded from sunset to sunrise at Walk-in Cave, Wood Buffalo National Park, Alberta, Canada, from mid-April to mid-October, during (a) 2011 and (b) 2012. Ambient temperature at sunset was obtained from Environment Canada for the Fort Smith Airport weather station. Nights with missing data (due to detector malfunction) are denoted with an ‘‘x’’. Note the y-axis scales differ between panels. chamber) on 10 June 2011 and 11 June 2012, entrance is a large opening located mid-way respectively. along a chamber, creating a T-shape with each Each time we entered the cave over the course arm extending ,25-m deep with a high ceiling. of the 3 y (6 visits: 10 and 11 September 2010; 16 We observed only 2 M. lucifugus, one of which May, 11 June, and 10 November 2011; and 10 was covered in ice droplets. June 2012), we observed bat carcasses on the cave floor. On 10 November 2012 we observed a Hibernaculum Conditions group of 6 M. lucifugus carcasses under a rock Temperature in Walk-in Cave was relatively fall in the Bat Chamber, 3 separate M. lucifugus stable throughout 2011 and 2012 (Fig. 4). The carcasses throughout the entrance hall and entrance chamber ranged between 1 and 26C middle chamber, and 2 E. fuscus carcasses in from June through October 2011, dropped the middle chamber, including 1 recently below 06C in mid-November 2011 (minimum deceased, face down in a shallow puddle of 24.16C), and returned to above 06C in mid- water. On 10 September 2010, a similar obser- April 2012. The bat chamber, where the majority vation was made of a recently dead adult of Myotis spp. were observed hibernating, female E. fuscus floating in the shallow cave fluctuated between 21.1 and 0.86C from 11 water. There were no clinical signs of white- November 2011 to 10 June 2012. nose syndrome (see Gargas and others 2009) on bats in the cave. Bat Activity at WB Pond and Surrounding Areas On 12 September 2010 we entered Peace Point Cave to investigate bat presence and general We recorded echolocation activity of both habitat suitability for hibernating bats. The resident and migratory species at WB Pond 282 NORTHWESTERN NATURALIST 95(3)

FIGURE 3. Mist net captures at the northern entrance of Walk-in Cave, Wood Buffalo National Park, Alberta, Canada, during 2011 (with the exception of captures on 12 May 2010, which occurred at the southern entrance of the sinkhole). Bars are stacked to represent total capture numbers; capture effort (netting minutes) is listed below each date (based on one 10- 3 2.6-m net). MYLU 5 Myotis lucifugus, MYSE 5 Myotis septentrionalis, EPFU 5 Eptesicus fuscus.

FIGURE 4. Hourly recordings of ambient temperature inside Walk-in Cave, Wood Buffalo National Park, Alberta, Canada. The highest concentration of hibernating bats was observed in the Bat Chamber. WINTER 2014 REIMER AND OTHERS:BAT ACTIVITY IN WOOD BUFFALO NP 283

FIGURE 5. (a) Resident and (b) migratory bat activity recorded during 2011 at WB pond, Wood Buffalo National Park, AB. Ambient temperature at sunset was obtained from Environment Canada at the Fort Smith Airport weather station. Nights with missing data (due to detector malfunction) are denoted with an ‘‘x’’. Note the y-axis scales differ between panels to best illustrate the data.

during 11 May to 7 October 2011 (n 5 166 nights). DISCUSSION The greatest bat activity was recorded during Species Diversity early August (Figure 5). We identified echoloca- tion files as Myotis spp. (n 5 23,002 files recorded We identified 5 bat species in WBNP: M. on 119 nights), E. fuscus (n 5 7624 files recorded lucifugus, M. septentrionalis, E. fuscus, L. cinereus, on 103 nights), L. cinereus (n 5 140 files recorded and L. borealis. As predicted, M. lucifugus, M. on 35 nights), and L. borealis (n 5 17 files septentrionalis, and E. fuscus were the most recorded on 8 nights). Ambient temperature at commonly detected bats, as illustrated by sunset ranged from 21.7 to 36.76C between 11 captures and echolocation recordings. This is June and 8 September 2011, and had a significant consistent with the wide distribution of M. effect on echolocation activity (linear regression, lucifugus (Fenton and Barclay 1980) and the 2 F1, 119 5 47.5, P , 0.001, R 5 0.29), with more presence of M. lucifugus and M. septentrionalis activity on warmer nights. We captured adult directly north (Reimer 2013; Wilson and others male and female M. lucifugus (n 5 188) and M. 2014) and south (Caceres and Pybus 1997; septentrionalis (n 5 58) throughout the summer, Vonhof and Hobson 2001; Grindal and others with the highest capture numbers from early 2011) of our study area. Despite the lack of August through early September when we records of E. fuscus in northeastern Alberta (300- began catching juveniles in addition to adults km south of WBNP; Grindal and others 2011), (Figure 6). We captured E. fuscus of both sexes captures of E. fuscus at Walk-in Cave, and and age classes (adult, juvenile), although far echolocation recordings at WB Pond were fewer (n 5 6) than Myotis spp. relatively common. 284 NORTHWESTERN NATURALIST 95(3)

FIGURE 6. Mist-net captures of bats in Wood Buffalo National Park at Pine Lake (14 September 2010), and sites within 3 km of Walk-in Cave (all other dates). Bars are stacked to represent total capture numbers; netting effort is listed below dates as min/m2. MYLU 5 Myotis lucifugus, MYSE 5 Myotis septentrionalis, EPFU 5 Eptesicus fuscus.

Two migratory species were detected in the the , L. noctivagans is reportedly study area: L. cinereus and L. borealis. Echoloca- relatively common (Grindal and others 2011). tion calls and captures of these migratory The absence of captures or echolocation record- species have been recorded directly south in ings in our study area suggests that this species northern Alberta (Patriquin 2004; Grindal and is either not present or uncommon; however, others 2011), and echolocation calls of L. cinereus due to similarities in echolocation calls between have been recorded as far north as , E. fuscus and L. noctivagans (Betts 1998), it is Northwest Territories (Wilson and others 2014) possible that some L. noctivagans calls were and southeastern Yukon (Slough and others misidentified as E. fuscus. In addition, our 2014). However, captures of either species have sampling was restricted geographically, and a yet to be reported in the Northwest Territories survey covering a larger area and different or Yukon. As these species roost solitarily and habitats may yet reveal the presence of L. often fly above the forest canopy (Kalcounis and noctivagans. others 1999; Menzel and others 2005), the lack of captures may be due to an emphasis on netting Seasonal Activity at ground level. Future studies should consider using stacked nets to gain height, and acoustic Echolocation activity at the mouth of Walk-in lures (Reyes and Szewczak 2010) to bring the Cave indicated that Myotis spp. and E. fuscus bats lower during flight. There are only 2 were active on or before 21 April 2011 and 2012, records of L. noctivagans farther north than our as activity was recorded immediately after study area: an acoustic recording and a photo- deploying the bat detectors. Activity continued graph of a live animal (Wilson and others 2014). through the end of September for both years The absence of this species in southwestern and ceased in mid-October in 2012. These Northwest Territories was also noted by Lausen timings are consistent with the onset and and others (2014). In northern Alberta, south of decline of activity at the nearby WB Pond, as WINTER 2014 REIMER AND OTHERS:BAT ACTIVITY IN WOOD BUFFALO NP 285 well as seasonal activity observed in other using torpor would provide energy savings for northern (Slough and Jung 2008; Talerico 2008; male M. lucifugus who do not bear the burden of Reimer 2013) and more southerly locations fetal development and lactation; however, (Davis and Hitchcock 1965; Anthony and others warmer temperatures are thought to be neces- 1981). In addition, an increase in Myotis spp. sary for spermatogenesis (Kurta and Kunz activity from early July to early September 1988). Perhaps, rather than a few males repeat- corresponds with the addition of flying young- edly using the cave as a summer day-roost, of-the-year, followed by the return of males and many individuals use it sporadically through- then females to hibernacula for autumn swarm- out the summer. ing (Hall and Brenner 1969; Thomas and others Juveniles of each resident species were cap- 1979; Schowalter 1980). These timings suggest tured in our study area, indicating that females that despite living at higher latitudes, Myotis reproduce in the area. This is not surprising spp. in WBNP may have a hibernation schedule given the close proximity of M. lucifugus and M. similar to those of species of Myotis occurring septentrionalis maternity colonies farther north farther south (Davis and Hitchcock 1965; Hall in the (Reimer 2013; Reimer and Brenner 1968). and Kaupas 2013), the presence of cabins used We performed extensive acoustic monitoring as roosts along Pine Lake, and tree roosting at the WB Pond during 2011 to record the opportunities across the study area. presence and seasonal distribution of species. L. cinereus was active in the area from mid-May to Hibernaculum Conditions early October, and L. borealis was active to a A majority of the Myotis spp. in Walk-in Cave lesser extent from mid-June to late August. The during September and November were ob- peak in activity of migratory species during late served roosting in the Bat Chamber. This is July and early August corresponds with au- consistent with where Lewis (1974–1975) re- tumn migration observed in more southern ported bats (he speculated that they were M. locations (Dalquest 1943; Findley and Jones lucifugus or E. fuscus) during the only winter 1964; Timm 1989; Cryan 2003; Baerwald and census performed prior to our work. He Barclay 2011). Resident bat species (Myotis spp. reported estimated populations of approximate- and E. fuscus) were more active in the area than ly 150 bats residing in the Bat Chamber while migratory species, and remained in the area our counts were lower (35–75 Myotis spp.). throughout the entire non-hibernating season. Although E. fuscus was not observed in the cave This is not surprising given the close proximity during our 2011 winter count, the capture of to hibernacula, and apparently suitable foraging individuals emerging from the cave during habitat. Resident bats also displayed an increase spring, the presence of carcasses throughout in activity with warmer ambient temperatures, the cave, and previous winter records of E. which has also been observed in several studies fuscus in the cave (Lewis 1974–1975) suggest farther south (for example, Menaker 1962; that some individuals hibernate in Walk-in Anthony and others 1981; Negraeff and Brig- Cave. We hypothesize that this species readily ham 1995; Erickson and West 2002; Dzal and roosts in crevices created by rock slabs in the Kunz 2012). cave. These rock slabs were common through- Myotis lucifugus, M. septentrionalis, and E. out the cave and were heard falling from the fuscus were all captured emerging from Walk- ceiling into the water at the bottom of the cave. in Cave during spring, and were captured This may explain the Big Brown Bats that frequently at the nearby WB Pond throughout appeared to have drowned. Torpid bats that the summer. Myotis lucifugus was the only fall into the water when a rock slab used for species observed using Walk-in Cave as a roosting falls would most likely drown before summer day roost. Other studies occasionally they could warm enough to swim. report a small number of M. lucifugus, typically Cave temperatures during the hibernation males, day-roosting at winter hibernacula dur- period (October to April) in the Bat Chamber ing the summer months (Davis and Hitchcock were warmer and more stable than the entrance 1965; Hall and Brenner 1968). Roosting in cooler chamber. These temperatures (21.1 to 0.86C) temperatures, such as in Walk-in Cave, and were colder than hibernacula temperatures at 286 NORTHWESTERN NATURALIST 95(3) lower latitudes (2.4 to 5.66C; Twente 1955; Beer areas of bat hibernation in North America. More and Richards 1956; Henshaw and Folk 1966; extensive mist-netting surveys in late July or Brack and Twente 1985; Davis and others 1999). early August that target higher flying species Because the thermal neutral zone of bats varies are likely to confirm the presence of migratory with latitude (Dunbar and Brigham 2010), bats species that we could only identify from hibernating in WBNP may have similar overall echolocation recordings. In addition, thermo- energy savings and expenditures during hiber- regulation studies investigating winter basal nation compared to southern conspecifics. As temperature of bats hibernating in Walk-in the Bat Chamber falls below freezing during the Cave would provide insight into both the effect winter months, further studies are needed to of colder cave temperatures on energy expen- assess basal temperature and torpor use of bats diture and the likelihood of establishment of P. in Walk-in Cave to investigate what additional destructans in the area. energetic expense the bats incur to remain above ambient temperature during periods ACKNOWLEDGMENTS below freezing, or what benefit they may have as a result of a lower basal temperature (slower We thank Julian Melnycky, Laura Kaupas, Brandon Klug, Dave Hobson, John McKinnon, Rhona Kindopp, metabolic rate) than bats farther south. Helen Panter, Jason Panter, and many of the Parks Evidence of infection by P. destructans, the Canada summer staff for assistance in the field. fungus that causes WNS (Minnis and Lindner Funding for this project was provided by the Alberta 2013), or clinical signs of disease (Gargas and Conservation Association and Bat Conservation Inter- others 2009) were not observed in Walk-in Cave. national to JPR, and the Natural Sciences and The hibernaculum temperatures were below the Engineering Research Council of Canada to RMRB. optimal growth conditions for the fungus (5 to Additional support was provided by Parks Canada 106C; Blehert and others 2009), although the and the Department of Environment and Natural fungus is able to grow at temperatures near Resources. We thank Greg Horne for providing a freezing. Further studies are required to deter- detailed description of Walk-in Cave for this manu- script; and we thank Paul Cryan, Tom Jung, and an mine the probability of WNS establishing under anonymous reviewer for suggestions that improved the conditions at this site. the paper. After investigating the Peace Point Cave, we suggest that it may be unsuitable for bat LITERATURE CITED hibernation. Given its shallowness and shape, we suspect temperature and humidity fluctua- ANTHONY ELP, STACK MH, KUNZ TH. 1981. Night tion would make this cave unsuitable for roosting and the nocturnal time budget of the hibernating bats, and that the 2 observed bats Little Brown Bat, Myotis lucifugus: Effects of in the cave would have moved to a deeper cave reproductive status, prey density, and environ- mental conditions. Oecologia 51:151–156 prior to the onset of winter. BAERWALD EF, BARCLAY RMR. 2011. Patterns of In conclusion, we found that WBNP is home activity and fatality of migratory bats at a wind to 2 migratory species (L. cinereus and L. borealis) energy facility in Alberta, Canada. Journal of and 3 resident species of bats (M. lucifugus, M. Wildlife Management 74:1103–1114. septentrionalis, and E. fuscus). With extensive BEER JR, RICHARDS AG. 1956. Hibernation of the Big industrial development occurring in northeast- Brown Bat. Journal of Mammalogy 37:31–41. ern Alberta, WBNP may provide important BETTS B. 1998. Effects of interindividual variation in intact habitat for bats, especially given the echolocation calls on identification of Big Brown presence of karst for hibernation, and wetlands, and Silver-haired Bats. Journal of Wildlife Man- riparian, and intact mixed-wood forests for agement 62:1003–1010. foraging and roosting. Given the vast region of BLEHERT DS, HICKS AC, BEHR M, METEYER CU, BERLOWSKI-ZIER BM, BUCKLES EL, COLEMAN JTH, karst topography and undeveloped habitat in DARLING SR, GARGAS A, NIVER R, OKONIEWSKI JC, WBNP, other hibernacula likely exist in the RUDD RJ, STONE WB. 2009. Bat white-nose syn- park. Further exploratory studies and efforts to drome: An emerging fungal pathogen? Science radio-track bats late in the summer may reveal 323:227. doi:10.1126/science.1163874. important hibernacula in this region, which BRACK JR V, TWENTE JW. 1985. The duration of the currently represents one of the most northern period of hibernation of three species of vesperti- WINTER 2014 REIMER AND OTHERS:BAT ACTIVITY IN WOOD BUFFALO NP 287

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